Original Investigation | Infectious Diseases

SARS-CoV-2 Transmission From People Without COVID-19 Symptoms

Michael A. Johansson, PhD; Talia M. Quandelacy, PhD, MPH; Sarah Kada, PhD; Pragati Venkata Prasad, MPH; Molly Steele, PhD, MPH; John T. Brooks, MD;
Rachel B. Slayton, PhD, MPH; Matthew Biggerstaff, ScD, MPH; Jay C. Butler, MD

Abstract Key Points

Question What proportion of
IMPORTANCE Severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2), the etiology of
coronavirus disease 2019 (COVID-19)
coronavirus disease 2019 (COVID-19), is readily transmitted person to person. Optimal control of
spread is associated with transmission
COVID-19 depends on directing resources and health messaging to mitigation efforts that are most
of severe acute respiratory syndrome
likely to prevent transmission, but the relative importance of such measures has been disputed.
coronavirus 2 (SARS-CoV-2) from
persons with no symptoms?
OBJECTIVE To assess the proportion of SARS-CoV-2 transmissions in the community that likely
occur from persons without symptoms. Findings In this decision analytical
model assessing multiple scenarios for
DESIGN, SETTING, AND PARTICIPANTS This decision analytical model assessed the relative the infectious period and the proportion
amount of transmission from presymptomatic, never symptomatic, and symptomatic individuals of transmission from individuals who
across a range of scenarios in which the proportion of transmission from people who never develop never have COVID-19 symptoms,
symptoms (ie, remain asymptomatic) and the infectious period were varied according to published transmission from asymptomatic
best estimates. For all estimates, data from a meta-analysis was used to set the incubation period at a individuals was estimated to account for
median of 5 days. The infectious period duration was maintained at 10 days, and peak infectiousness more than half of all transmission.
was varied between 3 and 7 days (−2 and +2 days relative to the median incubation period). The
Meaning The findings of this study
overall proportion of SARS-CoV-2 was varied between 0% and 70% to assess a wide range of
suggest that the identification and
possible proportions.
isolation of persons with symptomatic
COVID-19 alone will not control the
MAIN OUTCOMES AND MEASURES Level of transmission of SARS-CoV-2 from presymptomatic,
ongoing spread of SARS-CoV-2.
never symptomatic, and symptomatic individuals.

RESULTS The baseline assumptions for the model were that peak infectiousness occurred at the + Supplemental content
median of symptom onset and that 30% of individuals with infection never develop symptoms and Author affiliations and article information are
are 75% as infectious as those who do develop symptoms. Combined, these baseline assumptions listed at the end of this article.

imply that persons with infection who never develop symptoms may account for approximately 24%
of all transmission. In this base case, 59% of all transmission came from asymptomatic transmission,
comprising 35% from presymptomatic individuals and 24% from individuals who never develop
symptoms. Under a broad range of values for each of these assumptions, at least 50% of new SARS-
CoV-2 infections was estimated to have originated from exposure to individuals with infection but
without symptoms.

CONCLUSIONS AND RELEVANCE In this decision analytical model of multiple scenarios of

proportions of asymptomatic individuals with COVID-19 and infectious periods, transmission from
asymptomatic individuals was estimated to account for more than half of all transmissions. In
addition to identification and isolation of persons with symptomatic COVID-19, effective control of
spread will require reducing the risk of transmission from people with infection who do not have
symptoms. These findings suggest that measures such as wearing masks, hand hygiene, social
distancing, and strategic testing of people who are not ill will be foundational to slowing the spread
of COVID-19 until safe and effective vaccines are available and widely used.

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Open Access. This is an open access article distributed under the terms of the CC-BY License.

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Introduction
As severe acute respiratory syndrome coronavirus 2 (SARS-CoV-2), the novel coronavirus that causes
coronavirus disease 2019 (COVID-19), began to spread globally, it became apparent that the virus,
unlike the closely related SARS-CoV in the 2003 outbreak, could not be contained by symptom-
based screening alone. Asymptomatic and clinically mild infections were uncommon during the
2003 SARS-CoV outbreak, and there were no reported instances of transmission from persons
before the onset of symptoms.1 SARS-CoV-2 spread faster than SARS-CoV, and accumulating
evidence showed that SARS-CoV-2, unlike SARS-CoV, is transmitted from persons without
symptoms. However, measures to reduce transmission from individuals who do not have COVID-19
symptoms have become controversial and politicized and have likely had negative effects on the
economy and many societal activities. Optimal control of COVID-19 depends on directing resources
and health messaging to mitigation efforts that are most likely to prevent transmission. The relative
importance of mitigation measures that prevent transmission from persons without symptoms has
been disputed. Determining the proportion of SARS-CoV-2 transmission that occurs from persons
without symptoms is foundational to prioritizing control practices and policies.
Transmission by persons who are infected but do not have any symptoms can arise from 2
different infection states: presymptomatic individuals (who are infectious before developing
symptoms) and individuals who never experience symptoms (asymptomatic infections, which we
will refer to as never symptomatic). Early modeling studies of COVID-19 case data found that the
generation interval of SARS-CoV-2 was shorter than the serial interval, indicating that the average
time between 1 person being infected and that person infecting someone else was shorter than the
average time between 1 person developing symptoms and the person they infected developing
symptoms.2-5 This finding meant that the epidemic was growing faster than would be expected if
transmission were limited to the period of illness during which individuals were symptomatic. By the
time a second generation of individuals was developing symptoms, a third generation was already
being infected. Epidemiological data from early in the pandemic also suggested the possibility of
presymptomatic transmission,6,7 and laboratory studies confirmed that levels of viral RNA in
respiratory secretions were already high at the time of symptom onset.8-10
Asymptomatic SARS-CoV-2 transmission also occurs because of individuals with infection who
are never symptomatic (or who experience very mild or almost unrecognizable symptoms). The
proportion of individuals with infection who never have apparent symptoms is difficult to quantify
because it requires intensive prospective clinical sampling and symptom screening from a
representative sample of individuals with and without infection. Nonetheless, evidence from
household contact studies indicates that asymptomatic or very mild symptomatic infections
occur,11-14 and laboratory and epidemiological evidence suggests that individuals who never
develop symptoms may be as likely as individuals with symptoms to transmit SARS-CoV-2
to others.9,15,16

Methods
The Centers for Disease Control and Prevention determined that this decision analytical study, which
involved no enrollment of human subjects, did not require institutional review board approval. We
used a simple model to assess the proportion of transmission from presymptomatic (ie, infectious
before symptom onset), never symptomatic, and symptomatic individuals across a range of
scenarios in which we varied the timing of the infectious period to assess different contributions of
presymptomatic transmission and the proportion of transmission from individuals who never
develop symptoms (ie, remain asymptomatic).
For all estimates we used data from a meta-analysis of 8 studies from China to set the
incubation period at a median of 5 days with 95% of symptomatic individuals developing symptoms

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by day 12.17 Therefore the daily (t) probability of symptom onset (pso) for individuals who develop
symptoms was:

pso(t) = FLog−Normal(t,logmean = 1.63,logsd = 0.5).

To approximate a distribution of the infectious period, we made a baseline assumption that peak
infectiousness occurs on average at the same time as the median incubation period, such that
infectiousness begins prior to symptom onset (Table).9,12,14-16,18,20 We then assumed that
infectiousness (I) over time can be approximated by a γ density function and that the average person
is infectious for as long as approximately 10 days (ie, 98% of transmission happens within a 10-day
period)11:

I(t) = fγ(t,mode = 5,interval = 10).

For all estimates, we maintained the infectious period duration as 10 days, but varied the mode
between 3 and 7 days (−2 and +2 days relative to the median incubation period).
Uncertainty also remains about the proportion of individuals with infection who are never
symptomatic (pns) and the relative contribution of these infections to transmission (rns). Estimates of
pns range from single digits to more than 50%, many with potential biases related to the study
population (eg, age, prevalence of comorbidities) and the extent of long-term follow-up12-14,19,20
(Table). We made a baseline assumption that 30% of individuals with infection are never
symptomatic and then assessed higher or lower assumptions. We also made a baseline assumption
that individuals with asymptomatic infections are on average 75% as infectious as those with
symptomatic infections.9,15,16 Combined, these baseline assumptions imply that persons with
infection who never develop symptoms may account for approximately 24% of all transmission (T):

Tns = pns × rns / (pns × rns + [1 − pns]).

We varied this overall proportion, Tns, between 0% and 70% to assess a wide range of possible
proportions. The daily proportion of transmission from individuals after symptom onset (Ts) was
therefore:

Ts(t) = (1 − Tns) × pso(t) × I(t),

Table. Key Assumptions and Evidence Informing Those Assumptions

Source Evidence base Estimate or assumption

Assumptions for presymptomatic transmission
Peak infectiousness relative to onset, d
Casey et al, 202018 Range, 17 studies −3 to 1.2 d
Assumed baseline NA 0d
Assumed range NA −2 to 2 d
Assumptions for never symptomatic transmission
Proportion never symptomatic
Oran et al, 202012 Inferred range, 16 studies 30% to 45%
Buitrago-Garcia et al, 202014 Meta-estimate, 7 studies 26% to 37%
Davies et al, 202020 Age-dependent estimate, 6 studies 20% to 70%
Assumed baseline NA 30%
Relative infectiousness of individuals who never
have symptoms
Lee et al, 20209 303 patients, assessment of viral Approximately 100%
shedding
Chaw et al, 202015 1701 secondary contacts 40% to 140%
Mc Evoy et al, 202016 Inferred range, 6 studies 40% to 70%
Assumed baseline 75%
Overall proportion of individuals who never
have symptoms transmission
Assumed range NA 0% to 70%
Abbreviation: NA, not applicable.

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and the daily proportion of transmission from presymptomatic (Tps) individuals, ie, those who
develop symptoms but become infectious prior to symptom onset, is:

Tps(t) = 1 − Ts(t) − Tns.

We modified baseline assumptions to consider the relative importance of different levels of never
symptomatic and presymptomatic transmission. Code is available in the eAppendix in the
Supplement.

Statistical Analysis
All analyses were conducted in R version 4.0.1 (R Project for Statistical Computing). No statistical
testing was conducted, so no prespecified level of significance was set.

Figure 1. The Contribution of Asymptomatic Transmission Under Different Infection Profiles

Peak infectiousness: day 4 Presymptomatic: 43% Peak infectiousness: day 5 Presymptomatic: 35% Peak infectiousness: day 6 Presymptomatic: 27%
Never symptomatic: 24% Symptomatic: 33% Never symptomatic: 24% Symptomatic: 41% Never symptomatic: 24% Symptomatic: 49%

A Presymptomatic transmission with peak B Presymptomatic transmission with peak C Presymptomatic transmission with peak
infectiousness at day 4 infectiousness at day 5 infectiousness at day 6
0.008 0.008 0.008
Relative hourly infectiousness

Relative hourly infectiousness

Relative hourly infectiousness

0.004 0.004 0.004

0 0 0
0 5 10 15 0 5 10 15 0 5 10 15
Postinfection time, d Postinfection time, d Postinfection time, d

Peak infectiousness: day 5 Presymptomatic: 42% Peak infectiousness: day 5 Presymptomatic: 35% Peak infectiousness: day 5 Presymptomatic: 32%
Never symptomatic: 8% Symptomatic: 50% Never symptomatic: 24% Symptomatic: 41% Never symptomatic: 30% Symptomatic: 38%

D Transmission from individuals who are never E Transmission from individuals who are never F Transmission from individuals who are never
symptomatic (8%) symptomatic (24%) symptomatic (30%)
0.008 0.008 0.008
Relative hourly infectiousness

Relative hourly infectiousness

Relative hourly infectiousness

0.004 0.004 0.004

0 0 0
0 5 10 15 0 5 10 15 0 5 10 15
Postinfection time, d Postinfection time, d Postinfection time, d

The top curve in each panel represents the average relative hourly infectiousness, such
that while the lower curves change under different assumptions, the total hourly
infectiousness equals 1 in all cases. Within each curve, the colored area indicates the
proportion of transmission from each class of individuals. The portion attributed to
individuals with symptoms (light blue) can also be interpreted as the maximum
proportion of transmission that can be controlled by immediate isolation of all
symptomatic cases. Panels A, B, and C show different levels of presymptomatic
transmission. We calibrated infectiousness to peak at day 4 (A), 5 (B; median incubation
period), or 6 (C) days. Panels D, E, and F show different proportions of transmission from
individuals who are never symptomatic: 8% (C; eg, 10% never symptomatic and 75%
relative infectivity), 24% (D; baseline, 30% never symptomatic and 75% relative
infectivity), and 30% (E; eg, 30% never symptomatic and 100% relative infectivity).

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Results
Under baseline assumptions, approximately 59% of all transmission came from asymptomatic
transmission: 35% from presymptomatic individuals and 24% from individuals who are never
symptomatic (Figure 1). Because each component is uncertain, we assessed different timings of peak
infectiousness relative to illness onset and different proportions of transmission from individuals
who never have symptoms. Maintaining the 24% of transmission from individuals who never have
symptoms, but shifting peak infectiousness 1 day earlier (to day 4) increased presymptomatic
transmission to 43% and all asymptomatic transmission to 67% (Figure 1A). A later peak (ie, day 6)
decreased presymptomatic to 27% and all asymptomatic transmission to 51% (Figure 1C).
Holding the day of peak infectiousness constant at day 5 and decreasing the proportion of
transmission from individuals who are never symptomatic to 10% with a relative infectiousness of
75% (baseline assumption), the proportion of all transmission from those who are never
symptomatic decreased to 8%, presymptomatic transmission increased to 42%, and combined
asymptomatic transmission was 50% of all transmission (Figure 1D). In contrast, if the proportion of
those who ever develop symptoms was 30% and their relative infectiousness increased to 100%,
they contributed 30% of all transmission, presymptomatic transmission was 32%, and combined
asymptomatic transmission was 62% of all transmission (Figure 1F).
Uncertainty remains regarding the magnitude of both presymptomatic and never symptomatic
transmission. Therefore, we analyzed a wider range of each of these components, with peak
infectiousness varying between 2 days before (more presymptomatic transmission) to 2 days after
(less presymptomatic transmission) median symptom onset and with never symptomatic
transmission ranging from 0% to 70% (Figure 2). Under this broader range of scenarios, most
combined assumptions of peak infectiousness timing and transmission from individuals who never
have symptoms indicated that at least 50% of new SARS-CoV-2 infections likely originated from
individuals without symptoms at the time of transmission. If more than 30% of transmission was
from individuals who never have symptoms, total asymptomatic transmission was higher than 50%
with any value of peak infectiousness, up to 2 days after the median time of symptom onset. If peak
infectiousness was at any point approximately 6 hours before median symptom onset time, more
than 50% of transmission was from individuals without symptoms, regardless of the proportion from
those who never have symptoms. Even a very conservative assumption of peak infectiousness 2 days

Figure 2. Combined Transmission From Individuals Who Are Presymptomatic and Those Who Never
Have Symptoms

70
All asymptomatic
75% transmission (%)
100
60
Proportion of transmission from individuals

80

50
who are never symptomatic, %

60

40 40

20
30 Colors indicate the proportion of transmission due to
all individuals without symptoms at the time of
0
transmission, including presymptomatic transmission
20 (x-axis, the timing of peak infectiousness relative to
symptom onset) and transmission from individuals
who are never symptomatic (y-axis). For example,
10
peak infectiousness at the same time as median
% symptom onset (0 days difference) with 10% of
50
0 transmission from individuals who never have
–2 –1.5 –1.0 –0.5 0 0.5 1.0 1.5 2.0
symptoms would mean that approximately 51% of
Time from peak infectiousness to median symptom onset, d
transmission is from asymptomatic individuals.

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post–median onset and 0% never symptomatic transmission still resulted in more than 25% of
transmission from asymptomatic individuals.

Discussion
The findings presented here complement an earlier assessment21 and reinforce the importance of
asymptomatic transmission: across a range of plausible scenarios, at least 50% of transmission was
estimated to have occurred from persons without symptoms. This overall proportion of transmission
from presymptomatic and never symptomatic individuals is key to identifying mitigation measures
that may be able to control SARS-CoV-2. For example, if the reproduction number (R) in a given
setting is 2.0, then at least a 50% reduction in transmission is needed to drive the reproductive
number below 1.0. Given that in some settings R is likely much greater than 2 and more than half of
transmissions may come from individuals who are asymptomatic at the time of transmission,
effective control must mitigate transmission risk from people without symptoms.

Limitations
This study has limitations. First, we used a simplistic model to represent a complex phenomenon, ie,
the average infectiousness of SARS-CoV-2 infections over time. We used this model deliberately to
test assumptions about the timing of peak infectiousness and transmission among asymptomatic
individuals so that we could vary only these 2 critical parameters and assess their relative effects.
Therefore, these results lack quantitative precision, but they demonstrate the qualitative roles of
these 2 components and show that across broad ranges of possible assumptions, the finding that
asymptomatic transmission is a critical component of SARS-CoV-2 transmission dynamics remains
constant.
As discussed here, the exact proportions of presymptomatic and never symptomatic
transmission are not known. This also applies to the incubation period estimates, which are based on
individual exposure and onset windows that are difficult to observe with precision and therefore
include substantial uncertainty even when leveraging estimates across multiple studies. Moreover,
they likely vary substantially in different populations. For example, older individuals are more likely
than younger persons to experience symptoms,20 so in populations of older individuals, never
asymptomatic transmission may be less important. However, specific age groups are rarely
exclusively isolated from other age groups, so asymptomatic transmission risk is still important in
those groups and even more so in younger age groups, in which transmission may be even more
dominated by asymptomatic transmission.20
Real-world transmission dynamics are also not entirely dependent on the individual-level
dynamics of infectiousness over time. Now that COVID-19 is widely recognized, individuals with
COVID-19 symptoms are more likely to isolate themselves and further reduce the proportion of
transmission from symptomatic individuals, shifting a greater proportion of transmission to those
who do not have symptoms. In this sense, the estimates here represent the lower end of the
proportion of asymptomatic transmission in the presence of interventions to reduce symptomatic
transmission.

Conclusions
Under a range of assumptions of presymptomatic transmission and transmission from individuals
with infection who never develop symptoms, the model presented here estimated that more than
half of transmission comes from asymptomatic individuals. In the absence of effective and
widespread use of therapeutics or vaccines that can shorten or eliminate infectivity, successful
control of SARS-CoV-2 cannot rely solely on identifying and isolating symptomatic cases; even if
implemented effectively, this strategy would be insufficient. These findings suggest that effective

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control also requires reducing the risk of transmission from people with infection who do not have
symptoms. Measures such as mask wearing and social distancing empower individuals to protect
themselves and, if infected, to reduce risk to their communities.21 These measures can also be
supplemented by strategic testing of people who are not ill, such as those who have exposures to
known cases (eg, contact tracing) or are at high risk of exposing others (eg, congregate facility staff,
those with frequent contact with the public). Multiple measures that effectively address transmission
risk in the absence of symptoms are imperative to control SARS-CoV-2.

ARTICLE INFORMATION
Accepted for Publication: December 7, 2020.
Published: January 7, 2021. doi:10.1001/jamanetworkopen.2020.35057
Open Access: This is an open access article distributed under the terms of the CC-BY License. © 2021 Johansson
MA et al. JAMA Network Open.
Corresponding Author: Jay C. Butler, MD, Office of the Deputy Director for Infectious Diseases, US Centers for
Disease Control and Prevention, 1600 Clifton Rd, Mailstop H24-12, Atlanta, GA 30329 (jcb3@cdc.gov).
Author Affiliations: COVID-19 Response, US Centers for Disease Control and Prevention, Atlanta, Georgia
(Johansson, Quandelacy, Kada, Prasad, Steele, Brooks, Slayton, Biggerstaff); Office of the Deputy Directory for
Infectious Diseases, US Centers for Disease Control and Prevention, Atlanta, Georgia (Johansson, Slayton,
Biggerstaff, Butler).
Author Contributions: Dr Johansson had full access to all of the data in the study and takes responsibility for the
integrity of the data and the accuracy of the data analysis.
Concept and design: Johansson, Quandelacy, Kada, Brooks, Slayton, Butler.
Acquisition, analysis, or interpretation of data: All authors.
Drafting of the manuscript: Johansson, Quandelacy, Brooks, Biggerstaff, Butler.
Critical revision of the manuscript for important intellectual content: Johansson, Kada, Prasad, Steele, Brooks,
Slayton, Biggerstaff, Butler.
Statistical analysis: Johansson, Quandelacy, Kada.
Administrative, technical, or material support: Prasad, Steele, Brooks, Biggerstaff, Butler.
Supervision: Johansson, Butler.
Conflict of Interest Disclosures: None reported.
Funding/Support: This work was performed as part of the US Centers for Disease Control and Prevention’s
coronavirus disease 2019 response and was supported solely by federal base and response funding.
Role of the Funder/Sponsor: The funder had no role in the design and conduct of the study; collection,
management, analysis, and interpretation of the data; preparation, review, or approval of the manuscript; and
decision to submit the manuscript for publication.
Disclaimer: The findings and conclusions in this report are those of the authors and do not necessarily represent
the views of the Centers for Disease Control and Prevention.

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SUPPLEMENT.
eAppendix. Code for Analysis

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